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XIII. Genetics of Quantitative Traits

XIII. Genetics of Quantitative Traits

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ture. The planet’s carrying capacity is not unlimited, and environmentalconstraints

are ever increasing. Moreover, the balance of demographic power has shifted to

the developing world, where about 78% of human beings live. Poverty is taking

its toll, and more than 1 billion people today survive on less than a dollar a day.

Immediate measures must be undertaken to provide quick and reasonable relief to

this large segment of society.

About one-sixth of the world’s population live in the semi-arid tropics encompassing parts of Asia, Africa, and Latin America-the regions typified by limited

and erratic rainfall and poor soils. Pearl millet provides sustenance to a large proportion of poor people in these regions. It has the capacity to grow in some of the

poorest soils in chronically drought-prone regions. The need for genetic improvement of pearl millet cannot, therefore, be overemphasized. Although its importance as a research tool in cytogenetics and breeding has been recognized, its potential as an economic crop has not been fully realized. This poses a challenge for

cytogeneticists,breeders, agronomists, and biotechnologists.

Pearl millet is endowed with an efficient C, photosynthetic pathway, and it

responds well to fertilizers. Although it has a remarkable ability to grow on poor,

depleted soils, nitrogen deficiency is a major factor limiting grain production.

Therefore, genotypes with high-nitrogen-use efficiency should be produced. Fortunately, pearl millet responds extremely well to heterosis breeding. Utilization of

hybrid vigor will, therefore, be the most efficient means of increasing both grain

and forage production. If the vast pearl millet growing areas in Africa and Asia

could be planted to improved hybrids, grain production would increase phenomenally. Apomixis provides a unique tool for reaping the fruits of heterosis over an

extended period of time. If apomixis is transferred to hybrids with desired heterozygosity and superior gene combination, it can fix and help perpetuate heterosis, thereby obviating the need to produce hybrid seed year after year. Research in

this area will be very rewarding.

Developing a broad genetic base of hybrids is imperative to ensuring resistance

to future diseases. With the availability of cytoplasmic-genic male-sterile lines in

the mid- 1960s, several excellent hybrids were produced in India. Particularly

promising among these was HB 3, which, because of its high yields, became widely accepted throughout India in the early 1970s. Soon afterward, however, the hybrid became vulnerable to downy mildew caused by the fungus Sclemsporu

gruminicolu. The disease devastated the relatively genetically uniform hybrid

crop. An effective solution to such an eventuality is to produce genetically broadbased male-sterile lines using disease-resistantgenetic resources. Recently, several male-sterile lines have been developed at ICRISAT, and thnx of these (ICMA

91113, ICMA 91114, and ICMA 91115) provide not only reasonable yields but

also resistance to ergot, smut, and even downy mildew.

Pearl millet is an important source of dietary protein for a sizable portion of

those living in poverty in Africa and Asia. Therefore, the nutritional quality of the



grain, particularly its protein content and amino acid balance, needs to be improved. With genetic enrichment of the quantity and quality of its proteins, pearl

millet will be a more nutritional food source.

Cytogenetic manipulations have no doubt been instrumental in producing superior cultivars of pearl millet. An exciting recent development is the availability

of tools of modern biotechnology for crop improvement.The development and use

of molecular markers-random amplified polymorphic DNA (RAPDs) and restriction fragment length polymorphism (RFLPs)-are beginning to revolutionize

molecular mapping. For example, until recently, our knowledge of the inheritance

of downy mildew resistance was limited. Resistance was generally believed to be

monogenic dominant. However, molecular mapping has demonstrated that many

genes contribute to downy mildew resistance and that these genes are scattered

throughout the host genome. The use of DNA markers could help identify desired

genotypes more precisely and hence assist in adopting appropriate breeding strategy for pearl millet.

Pearl millet provides unlimited opportunities for both basic and applied research. With further cytogenetic manipulation and marker-assisted selection, combined with the exploitation of recent advances in biotechnological research, pearl

millet may emerge as a leading economic crop that plays an ever-increasing role

in the welfare of those living in poverty, particularly in the semi-arid tropics of the



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Parviz N. Soltanpour,' Greg W. Johnson,2 Stephen M. W ~ r k m a n , ~

J. Benton Jones, Jr.: and Robert 0. Miller'

'Department of Soil and Crop Sciences

Colorado State University

Fort Collins, Colorado 80523

2Matheson Gas Products

Longmont, Colorado 80501

'Analytical Technologies, Inc.

Fort Collins, Colorado 80524

'+Macro-MicroAnalytical Services

Athens, Georgia 30607

I. Introduction

11. ICP-AES and ICP-MS Instrumentation

A. ICP Generation

B. Properties of ICP

C. Sample Introduction Systems

111. Spectrometers

A. ICP-Atomic Emission Spectrometry

B. ICP-Mass Spectrometry

n! Analytical Capabilities

A. Selection of Wavelength

B. Selection of Isotope

C. ICP-AES Detection Limits

D. ICP-MS Detection Limits

v. ICP-AES Interferences

A. Solute Vaporization

B. Ionization

C. Unwanted Radiation

D. Correction for Interferences (ICP-ms)

VI. ICP-MS Interferences

A. Solids Deposition on Sampler and Skimmer Cones


Advrmrm in Agrorm~y,Volume 64

Copynght 0 1998 by Academic Press. All rights of reproduction in any form reserved.





B. NonspectroscopicInterferences

C. Mass Discrimination

D. Unwanted Ions

E. Methods of Correction for Interferences(ICP-MS)

VII. Practical Applications

A. Grinding Soil Samples

B. Obtaining Soil Extracts

C. Digestion of Organic Matter and Dissolution of Silicates for Total

Elemental Analysis

D. Analysis of Soil Extracts and Digests

E. Determination of Trace Levels of As, Se, and Hg Using the HydrideMercury Vapor Generator

VIII. Quality Control Methods





The application of inductively coupled plasma-atomic emission spectrometry

(ICP-AES) to the analysis of soil was reviewed in 1982 and again in 1996 with inclusion of ICP-mass spectrometry (ICP-MS) (Soltanpour et al., 1982, 1996). In

this review we treat ICP-MS more comprehensively and include a table for isotopes of elements (see Section 1V.B.) and an example for Ca, Fe, Ni, Zn, and Pb

isotope selection for plant-tissue analysis (Appendix 1).

New developments in ICP-AES include suspension-nebulization analysis of

clays (Laird et al., 1991); interfacing ICP spectrometers with flow-injection analyzers for automatic dilution, calibration, separation, concentration, standard additions, and other operations (Greenfield, 1983; LaFerniere et al., 1985); interfacing ICP-AES with liquid chromatographs for concentration and speciation of

elements (Roychowdhury and Koropchack, 1990); using high-salt nebulizers to

prevent clogging of nebulizers (Legere and Burgener, 1985): successfully using

concentration and reduction of spectral interference techniques such as chelation-solvent extraction (Huang and Wai, 1986; Bradford and Bakhtar, 1991); using computer programs such as orthogonal polynomials (Hassan and Loux, 1990),

simplex optimization (Belchamber et al., 1986), and that recommended by Taylor

and Schutyser (1986) to optimize spectrometer operating conditions and automatic correction for spectral interferences; and compiling ICP emission lines still in

progress (McLaren and Berman, 1985; Boumans, 1984; Parsons et al., 1980).

The ICP-MS method of analysis has been developed over the last 15 years.

Houk et al. (198 1) showed suprathermal ionization in an ICP Ar plasma. Within

the last 10 years the method has been applied to routine analytical concentration

determinations. Several review articles document the ICP-MS developmental

milestones (Beauchemin, 1989; Hieftje and Vickers, 1989; Douglas, 1989; Houk

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XIII. Genetics of Quantitative Traits

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